Distribution of clonal growth forms in wetlands

Clonal multiplication is a predominant type of reproduction in wetland species. However, both wetlands and plant organs of clonal growth are diverse, thus due to different stress factors operating in various wetlands preponderance of plants with specific clonal growth organs (CGOs) can be expected. To test this hypothesis the CGO spectra of wetland communities of the Netherlands were analysed, including a bog, a fen, heathland, a floodplain, river beds, fresh water pools, open salt water and a salt marsh. Moreover, it was evaluated whether different CGOs are characterised by different functional traits (shoot cyclicity, persistence of connections between ramets, number of offspring produced per year and lateral spread per year) in wetland species. Data on types of CGO, i.e., epigeogenous and hypogeogenous rhizomes, fragments and budding plants, stolons, tubers and bulbs, root-splitters, root-sprouters and special adaptations (turions) as well as their functional traits, were taken from the CLO-PLA 3 database. CGO spectra of wetland communities were analysed using two methods: comparison of observed vs. expected CGO spectra based on the presence/absence data and multivariate analysis (CCA) for inter-community differences considering species frequency. Moreover, relationships between CGOs and their functional traits were tested using multidimensional contingency tables. Apart from 26% of non-clonal species, the majority of wetland species was rhizomatous (51%). Other types of CGO were represented in less than 10% of species and root-derived CGOs were underrepresented (<2%) in comparison with terrestrial habitats. Among communities, fresh water pools and open salt water hosted higher proportion of species with fragments (∼10%) and turions (∼30%). Multivariate analysis divided wetland communities along the disturbance and hydric (water) gradients. Highly disturbed communities (salt marshes) were characterised by non-clonal species and species with root-derived CGOs. Aquatic communities (fresh water pools and open salt water) hosted species with the ability to spread by fragmentation and turions, contrary to permanently wet communities (bog and wet heathland) with the prevalence of species with epigeogenous rhizomes. It was also confirmed that the CGOs of wetland species differed in their traits. The most important functional trait characterising individual CGOs in the wetland flora was the degree of lateral spread (explained variability: 53%) followed by duration of persistence of connections between ramets (explained variability: 74%), which is in accordance with earlier distinguished strategies of clonal growth: integrator/splitter and spreading/non-spreading clones.     

Clonal splitters and integrators in harsh environments of the Trans-Himalaya

Individuals of clonal plants consist of physically and physiologically connected ramets. In splitters, they are integrated for a time shorter than ramet generation time (i.e. the time it takes to produce the first offspring ramet), whereas in integrators connections between ramets persist for a longer time. It has been predicted that integrators should prevail in stressful environments, such as habitats poor in nutrients, whereas splitters are expected to dominate in benign habitats, such as fertile areas with a moderate climate. I tested these predictions in four dry mountain areas of the Trans-Himalaya, in high altitudes subjected to multiple stresses. In accordance with the expectations I found that clonal plants with integrated ramets reach higher mean and maximum altitudes than splitters. Integrators were over-represented in nutrient-poor habitats, such as dry semi-deserts, sandy steppes and in subnival habitats, whereas splitters preferentially colonised mesic habitats, saline sites and wetlands. While there was no difference in the representation of splitters and integrators in habitats with an unstable surface, such as screes, dunes and water bodies, fully integrated clonal plants preferred very stable environments, such as banks of streams covered by closed-canopy vegetation. Most relationships between clonal integration and environmental factors were explainable by the phylogenetic relationship between the species, only the significant preference of splitters for shaded environments persisted in phylogenetically corrected analysis. The results indicate that clonal integration belongs to a set of evolutionarily conservative plant traits, usually shared by related species. Consequently, the adaptive value of clonal integration in individual habitats remains questionable.     

Spatial distribution pattern of a clonal species: effects of differential production of clonal and sexual offspring

The spatial distribution patterns of genets and ramets within populations are expected to change as a function of the frequency with which clonal species recruit different types of offspring (sexual and clonal). We used an integrated approach to study the spatial arrangement of clonal plants by combining molecular and ecological data using Opuntia microdasys as a study system. The species is able to produce two types of clonal (plantlets and cladodes) and one type of sexual (seeds) offspring. Additionally it is found in three habitats that cause differences in the ability of each type of offspring to establish. In 2007, all individuals in the three habitats (162 in BH = bajada, 264 in IDH = hill-piedmont, and 136 at HPH = interdunes) were tagged and mapped. Amplified inter-simple sequence repeats (ISSR’s) were used to determine the multilocus genotype and relatedness of each individual ramet using 120 polymorphic bands (104 in BH, 128 in HPH and 180 in IDH). The spatial distribution pattern of genets and ramets was analyzed with the Hopkins test and spatial autocorrelation analysis. For all habitats we found that O. microdasys displayed a spatial distribution characterized by clumps of aggregated ramets, but habitats differed in the number of genets present. As for other clonal species a strong positive spatial autocorrelation exists within 20 m, although all analyses suggest that adjacent ramets are genetically less related to each other or belong to different genets, that is, ramets of different genets are intermingled. The spatial arrangement of genets and ramets in O. microdasys between habitats closely matches the frequency of establishment of each type of offspring (e.g. the more clonal areas are clumped groups of related individuals). These results confirm that in two habitats (BH and IDH) clonal recruitment had been more common than in the other habitat (HPH).     

Small-scale heterogeneity in soil quality influences photosynthetic efficiency and habitat selection in a clonal plant

BACKGROUND AND AIMS: In clonal plants, internode connections allow translocation of photosynthates, water, nutrients and other substances among ramets. Clonal plants form large systems that are likely to experience small-scale spatial heterogeneity. Physiological and morphological responses of Fragaria vesca to small-scale heterogeneity in soil quality were investigated, together with how such heterogeneity influences the placement of ramets. As a result of their own activities plants may modify the suitability of their habitats over time. However, most experiments on habitat selection by clonal plants have not generally considered time as an important variable. In the present study, how the foraging behaviour of clonal plants may change over time was also investigated. METHODS: In a complex of environments with different heterogeneity, plant performance was determined in terms of biomass, ramet production and photosynthetic activity. To identify habitat selection, the number of ramets produced and patch where they rooted were monitored. KEY RESULTS: Parent ramets in heterogeneous environments showed significantly higher maximum and effective quantum yields of photosystem II than parents in homogeneous environments. Parents in heterogeneous environments also showed significantly higher investment in photosynthetic biomass and stolon/total biomass, produced longer stolons, and had higher mean leaf size than parents in homogeneous environments. Total biomass and number of offspring ramets were similar in both environments. However, plants in homogeneous environments showed random allocation of offspring ramets to surrounding patches, whereas plants in heterogeneous environments showed preferential allocation of offspring to higher-quality patches. CONCLUSIONS: The results suggest that F. vesca employs physiological and morphological strategies to enable efficient resource foraging in heterogeneous environments and demonstrate the benefits of physiological integration in terms of photosynthetic efficiency. The findings indicate that short-term responses cannot be directly extrapolated to the longer term principally because preferential colonization of high-quality patches means that these patches eventually show reduced quality. This highlights the importance of considering the time factor in experiments examining responses of clonal plants to heterogeneity.     

扎龙湿地不同生境芦苇分株的形态特征及生物量分配的异速关系

不同环境条件下的植物个体可以表现出形态特征的分异和物质分配的权衡与调整。采用大样本抽样调查与统计分析方法,比较研究扎龙湿地不同生境芦苇(Phragmites Australis)生殖株和营养株的形态特征以及生物量分配的异速关系。结果表明：在9月末,盐碱生境、旱生生境、湿生生境和水生生境芦苇分株的生长表现出较大的生态可塑性,株高和株重均以盐碱生境最小,水生生境最大,最大值与最小值的比值分别为1.3—3.3和1.8—5.1,分株生长在种群间的变异度高于种群内,与营养株相比,生殖株的变异度较低;分株的支持分配与生产分配的比值为1.8—4.2,生产分配以盐碱生境最高,以水生生境最低,而支持分配和生殖分配表现与生产分配相反的序位;生殖株的花序长和花序重与株高间呈直线函数形式增长,株高和株重低于种群平均值的20%和35%的分株不进行有性生殖;叶重、叶鞘和茎重以及分株重与株高间呈幂函数形式的异速生长关系。植物通过改变个体的形态特征以及调整构件间生物量分配适应不同环境,而受遗传因素控制的构件间生长关系却相对稳定。     

Effects of the loss of clonal integration on four sedges that differ in ramet aggregation

Although clonal growth is a dominant mode of plant growth in wetlands, the importance of clonal integration, resource sharing among ramets, to individual ramet generations (mother and daughter) and entire clones of coexisting species has not been well investigated. This study evaluated the significance of clonal integration in four sedge species of varying ramet aggregations, from clump-forming species (Clumpers –Carex sterilis, Eleocharis rostellata), with tightly aggregated ramets (rhizomes<1cm), to runner species (Runners –Schoenoplectus acutus, Cladium mariscoides), with loosely aggregated ramets. We manipulated clonal integration by either severing connections between target mother and daughter ramets or leaving connections intact, and then planted them in an intact neighborhood of a fen in Michigan, USA. We measured growth parameters of original and newly produced ramets over two growing seasons and conducted a final biomass harvest, to address four hypotheses. First, we expected integrated clones to accumulate more biomass than severed clones. However, final clone-level biomass and ramet production were the same for both treatments in all species although severing initially stimulated ramet production by Schoenoplectus and produced a more compact ramet aggregation in Cladium. Second, we hypothesized that mother ramets would experience a cost of integration, through reduced ramet or biomass production, while daughters would experience a benefit, through increased resource availability from mothers. Mother ramets of Cladium suffered a cost from integration, while Schoenoplectus mothers suffered a slight cost and Carex daughters saw a slight benefit. Finally, we hypothesized that integration would be more active in runner species than in clumper species. Indeed, we documented more active integration in runners than clumpers, but none of the study species were dependent upon integration for growth or survival once daughter ramets were established with their own roots and shoots. This study demonstrates that integration between established ramets may not be the most important advantage to clonal growth in this wetland field site. The loss of integration elicited varied responses among coexisting species in their natural habitat, somewhat but not completely related to their growth form, suggesting that a combination of plant life history traits contributes to the dependence upon clonal integration among established ramets of clonal species.     

Clonal integration in Fragaria vesca growing in metal-polluted soils: parents face penalties for establishing their offspring in unsuitable environments

Clonal plants often establish descendent ramets in sites with contrasting presence of favourable and unfavourable factors. Connections between ramets allow translocation of essential resources from established ramets to developing ramets and, as consequence, integration confers net benefits to ramets growing under unfavourable conditions. Therefore, integrated ramets may survive in habitat patches that would be lethal to independent ramets or non-clonal plants. This experiment aimed to investigate the physiological and morphological responses of the clonal plant Fragaria vesca growing in heterogeneous substrate with patches of contrasting quality (i.e. uncontaminated or heavy-metal-contaminated). We observed that parents reduced their photosynthetic efficiencies and growth as consequence of maintaining their offspring. This cost did not affect survival of the parents. Physiological integration brings about benefits to offspring ramets growing both at uncontaminated and heavy-metal-contaminated soils. The benefits of integration were detected in both physiological and morphological traits, enhancing the survivorship of offspring ramets in the Cu-polluted soils. We conclude that integration improves the performance of developing ramets of F. vesca growing in heavy-metal-contaminated habitats, allowing clone systems to overcome the establishment risks and maintain their presence in these less favourable sites.     

Physiological integration ameliorates effects of serpentine soils in the clonal herb Fragaria vesca

Small-scale heterogeneity in soil characteristics and the facility of clonal systems to spread may lead to situations where parent ramets in favourable microhabitats are connected to offspring in stressful conditions. Clonal plants are physiologically integrated if connections among ramets allow transport of resources. Thus, ramets in favourable habitats may provide support to developing or stressed ramets. We examined effects of integration in Fragaria vesca growing in patches of contrasting quality (potting compost vs serpentine soil). Serpentine soil was used to create unfavourable growing conditions. We assessed whether survival, biomass and photosynthetic efficiency (estimated by fluorescence and reflectance) of parents and offspring were affected by integration and soil quality. Integration increased photochemical efficiencies of parents but more consistently in parents with offspring growing in serpentine soils. We suggest that the assimilate demand from offspring enhanced the photosynthetic efficiency of parents by a mechanism of feedback regulation. This result extends the concept of physiological integration in clonal plants to include photochemical responses. Connected parents also showed significantly higher biomass than disconnected parents. In our system, integration did not entail costs for the whole clone in terms of biomass. Integration also improves the survival, growth and photochemical efficiency of developing ramets, suggesting that integration represents a mechanism for increasing survival in stressful habitats, as the serpentine soils.     

Ephemeral clonal integration in Calathea marantifolia (Marantaceae): Evidence of diminished integration over time

A major advantage of clonal growth forms is the intergenerational transfer of resources through vascular connections (clonal integration). Connections linking ramets can be persistent or ephemeral. For species with ephemeral connections, whether the extent of clonal integration changes over time is unclear. To address this issue, we tracked water movement using an isotopic label and assessed the demographic performance of parent and offspring ramets over time in a severing experiment. Our study system was the understory herb Calathea marantifolia, which has parent ramets that produce vegetative bulbils (clonal offspring) that pass through distinct pre- and post-rooting stages. Little water was transported between parents and offspring, and the direction of movement was primarily from parent to pre-rooting offspring. Anatomical observations of inter-ramet connections showed that vascular bundles were twice as abundant in parent stems compared to inter-ramet connections. Severing inter-ramet connections reduced the growth of offspring ramets but not parents. Survival of pre-rooting offspring was reduced by 10% due to severing, but post-rooting offspring were not affected. Our results suggest that offspring ramets of C. marantifolia are weaned from their parent as they progress from pre- to post-rooting stages.     

The significance of rhizome connection of semi-shrub Hedysarum laeve in an Inner Mongolian dune,China

Hedysarum laeve, a rhizomatous clonal semi-shrub, commonly dominates the inland dunes in semiarid areas of northern China. This species propagates vegetatively by extension of horizontal woody rhizomes resulting in programmed reiteration of apical and/or axillary meristems. In this study, the plants were experimentally manipulated by cutting rhizome connections and ¹⁴C-labelling techniques were employed to investigate the ecological significance of rhizome connections within the H. laeve clone. Severance of rhizome connections had a great effect on the performance of young ramets within a clone. Young ramets severed from their parent ramets experienced a significant reduction both in ramet growth and vegetative propagation, as compared with the intact young ramets. Within clonal fragments, consisting of three interconnected ramets including a mother ramet, a daughter ramet and a granddaughter ramet, ¹⁴C-photosynthates from the fed leaves of mother ramets were acropetally transported to all clonal component parts. The ¹⁴C-photosynthate translocation within the clonal fragment provides evidence that the young ramets were supported by their parent ramets. Our results suggest that the woody rhizome connections among the interconnected ramets are ecologically and strategically important for the species to grow in the sand dune habitat.     

Clonal integration supports the expansion from terrestrial to aquatic environments of the amphibious stoloniferous herb Alternanthera philoxeroides

Effects of clonal integration on land plants have been extensively studied, but little is known about the role in amphibious plants that expand from terrestrial to aquatic conditions. We simulated expansion from terrestrial to aquatic habitats in the amphibious stoloniferous alien invasive alligator weed ( Alternanthera philoxeroides ) by growing basal ramets of clonal fragments in soils connected (allowing integration) or disconnected (preventing integration) to the apical ramets of the same fragments submerged in water to a depth of 0, 5, 10 or 15 cm. Clonal integration significantly increased growth and clonal reproduction of the apical ramets, but decreased both of these characteristics in basal ramets. Consequently, integration did not affect the performance of whole clonal fragments. We propose that alligator weed possesses a double-edged mechanism during population expansion: apical ramets in aquatic habitats can increase growth through connected basal parts in terrestrial habitats; however, once stolon connections with apical ramets are lost by external disturbance, the basal ramets in terrestrial habitats increase stolon and ramet production for rapid spreading. This may contribute greatly to the invasiveness of alligator weed and also make it very adaptable to habitats with heavy disturbance and/or highly heterogeneous resource supply.     

WATER SHARING AMONG RAMETS IN A DESERT POPULATION OF DISTICHLIS SPICATA (POACEAE)

Clonal growth enables plants to transport resources among separately rooted but connected ramets, a potential advantage in patchy or unpredictable habitats. Nevertheless, clonal plants are relatively scarce in deserts. To test whether clonal integration of water relations can increase plant performance under natural conditions in a desert species, water movement was traced and connection among ramets was manipulated in the rhizomatous grass Distichlis spicata in Death Valley, California. To examine potential costs of clonal growth form, connections were mapped and analyzed for dry mass and nitrogen content. Movement of dye showed potential transport of water among five ramets up to 1.4 m apart. Severance of connecting rhizomes increased mortality and decreased water potential of individual ramets within 36 hr, indicating that water sharing among ramets could be of significant benefit. However, plants had a high investment of mass and nitrogen in underground organs, which might be a cost of clonal growth associated with desert environments.     

To share or not to share: clonal integration in a submerged macrophyte in response to light stress

The ability of clonal plant species to share resources has been studied in many experiments. The submerged macrophyte Potamogeton perfoliatus produces interconnected ramets within short time intervals and hence may or may not share resources with ramets growing in less favourable microhabitats. From a genet point of view, sharing with ramets growing under less favourable conditions might not be an optimal strategy when photosynthates could be used to establish other ramets growing under more favourable conditions. To analyse the plasticity in clonal integration of P. perfoliatus, we set up a factorial aquaria experiment with unshaded or shaded recipient ramets (offspring), which were connected to or separated from donor ramets (parents). Increased biomass production of offspring in parent–offspring systems compared with severed offspring in both light and shade showed that ramets share resources through clonal integration. The relative translocation to the first- and second-offspring generation was influenced by habitat quality: If first-offspring ramets grew in a shaded microhabitat, second-offspring ramets clearly profited. This may be at least partially because of the fact that resources are shifted from first-offspring to second-offspring ramets, indicating controlled senescence of the first-offspring. This complex sharing behaviour might be relevant when plants produce ramets within a dense patch of macrophytes, where support of a shaded ramet might not pay off.     

Physiological integration helps a clonal macrophyte spread into competitive environments and coexist with other species

Physiological integration may help clonal macrophytes invade or escape from existing communities. No studies have tested the above hypothesis in aquatic plants. In an outdoor pond experiment, we subjected clonal fragments of the submerged macrophyte Vallisneria spiralis L. to heterogeneous environments in which V. spiralis spread from bare habitats towards vegetated habitats occupied by Myriophyllum spicatum L. or V. spiralis spread from vegetated habitats towards bare habitats. V. spiralis stolons between ramets in bare habitats and in vegetated habitats were either intact or severed. We investigated the habitat selection of V. spiralis by examining the allocation of biomass and ramets to heterogeneous habitats during its vegetative spread phase. Results showed that the stolon connection had different effects on the habitat selection of V. spiralis with regard to invasion and escape. When V. spiralis spread from bare to vegetated habitats, in comparison to severing the stolon, the stolon connection eventually facilitated a 49% increase in biomass and a 27% increase in number of ramets allocated to vegetated habitats. However, when V. spiralis spread from vegetated to bare habitats, biomass and ramets allocated to bare habitats were not significantly changed by the stolon connection (only a 5% increase in biomass and a 6% increase in number of ramets). These results indicate that clonal integration facilitated V. spiralis not to escape from but invade into vegetated habitats. The study provides evidence that physiological integration is important for survival and tolerance of ramets in competitively stressful environments and can help clonal macrophytes coexist with other species.     

Ramet production byMallotus japonicus, a common pioneer tree in temperate Japan

To clarify the characteristics of clonal growth ofMallotus japonicus, we excavated all roots of this species in a plot and ascertained the connections between shoots. All shoots examined were ramets that originated from roots. Young ramets grew in shady sites as well as sunnier sites, indicating that ramet production was not dependent on local light conditions.     

Ramet phenology and clonal architectures of the clonal sedge <Emphasis Type="Italic">Schoenoplectus americanus</Emphasis> (Pers.) Volk. ex Schinz &; R. Keller

The clonal plant Schoenoplectus americanus shows variable belowground clonal architecture as a result of producing two types of ramets: those with very long rhizomes (long rhizome ramet, LRR) and those with very short ones (short rhizome ramet, SRR). In a previous study we demonstrated that the two types of ramets are functionally specialised. The production of SRRs results in the formation of consolidated clonal patches with densely packed shoots, while the production of LRRs results in a more diffuse network of connected rhizomes with widely spaced shoots. We hypothesised that the two types of ramets would be produced at different times during the growing season because of their functional differences. The production of LRRs throughout the growing season would enable the species to continuously explore new habitats while the production of SRRs early in the growing season would enable the species to occupy and consolidate resources in available open patches. We evaluated this hypothesis through field observations in different communities with S. americanus and indeed found that SRRs were produced early in the growing season while LRRs tended to be produced over an extended period of time. Plants in high-quality environments (i.e. higher light conditions) produced more SRRs, and these were formed early in the growing season. In contrast, plants in low-quality environments produced more LRRs, and these were formed continuously over the growing season. We also observed that the shoot longevity was greater for SRR. In high-quality patches, the production of the lower cost SRRs results in a more rapid occupancy of open spaces; in lower quality patches, the production of LRRs throughout the growing season enables plants to explore the immediate environment for higher quality patches.     

Clonal integration ameliorates the carbon accumulation capacity of a stoloniferous herb,Glechoma longituba,growing in heterogenous light conditions by facilitating nitrogen assimilation in the rhizosphere

Background and Aims Enhanced availability of photosynthates increases nitrogen (N) mineralization and nitrification in the rhizosphere via rhizodeposition from plant roots. Under heterogeneous light conditions, photosynthates supplied by exposed ramets may promote N assimilation in the rhizosphere of shaded, connected ramets. This study was conducted to test this hypothesis.Methods Clonal fragments of the stoloniferous herb Glechoma longituba with two successive ramets were selected. Mother ramets were subjected to full sunlight and offspring ramets were subjected to 80 % shading, and the stolon between the two successive ramets was either severed or left intact. Measurements were taken of photosynthetic and growth parameters. The turnover of available soil N was determined together with the compostion of the rhizosphere microbial community.Key Results The microbial community composition in the rhizosphere of shaded offspring ramets was significantly altered by clonal integration. Positive effects of clonal integration were observed on NAGase activity, net soil N mineralization rate and net soil N nitrification rate. Increased leaf N and chlorophyll content as well as leaf N allocation to the photosynthetic machinery improved the photosynthetic capability of shaded offspring ramets when the stolon was left intact. Clonal integration improved the growth performance of shaded, connected offspring ramets and whole clonal fragments without any cost to the exposed mother ramets.Conclusions Considerable differences in microbial community composition caused by clonal integration may facilitate N assimilation in the rhizosphere of shaded offspring ramets. Increased N content in the photosynthetic machinery may allow pre-acclimation to high light conditions for shaded offspring ramets, thus promoting opportunistic light capture. In accordance with the theory of the division of labour, it is suggested that clonal integration may ameliorate the carbon assimilation capacity of clonal plants, thus improving their fitness in temporally and spatially heterogeneous habitats.     

On the ecological and evolutionary significance of storage in clonal plants

Environmental heterogeneity has received wide attention in clonal plant research over the last decade. Most studies have focussed on the effects of spatial variation in environmental conditions on the performance of ramets and genets, while the effects of temporal heterogeneity have only occasionally been studied. As a consequence, our current knowledge about functional responses of clonal plants to habitat patchiness is biased towards spatial aspects of environmental heterogeneity. Nevertheless, temporal changes in biotic and abiotic conditions do occur in most natural habitats, and they are very likely to affect plant growth and performance, and to create positive selection pressures on traits that can buffer plants against unfavorable consequences of this variability. Storage of resources is a widespread phenomenon in clonal plant species. Typical clonal structures such as stolons, rhizomes and hibernacles serve as storage organs. However, the ecologic significance of storage in clonal plant structures remains partly unclear. We suggest that the lack of understanding with respect to resource storage in clonal plants be related to our poor knowledge of ecologic implications of temporal habitat heterogeneity in natural environments. Resource storage can be understood as a safety measure against temporal changes in the growing conditions of plants. This paper summarizes existing information about the ecologic relevance of storage in clonal plants and it tries to develop a framework for further investigation of resource storage as a strategy to enhance the performance of clonal plants in temporally variable environments.     

Exploitation of patchy soil water resources by the clonal vine <Emphasis Type="Italic">Ficus tikoua</Emphasis> in karst habitats of southwestern China

The karst habitats of southwestern China are characterized by a highly heterogeneous distribution of water resources. We hypothesized that the clonal integration between connected ramets of the clonal vine Ficus tikoua was an important adaptive strategy to the patchy distribution of water resources in these habitats. We grew ramet pairs (each consisting of a parent and an offspring ramet) in both homogeneously and heterogeneously watered conditions. The offspring ramets were well-watered, whereas their connected parent ramets were randomly assigned to four water treatments: well-watered, mild water stress, moderate water stress, and severe water stress. Increasing water stress decreased leaf water potential, relative water content, net assimilation rate, maximum quantum yield of PSII (F _v/F _m), and biomass of the parent ramets. Subjecting the parents to water stress significantly increased root biomass and root mass ratio (RMR) of their offspring ramets. Exploitation of plentiful water resources through the increased adventitious roots connected to another soil patch permitted the complete restoration of water relations and photosynthetic capacity of offspring ramets after an initial depression. Water relations and gas exchange of the parents were not affected by the water supply to their connected offspring ramets, suggesting that offspring ramets hardly exported water to the stressed parents. However, net assimilation rate and proline content of the offspring ramets increased when they were connected to water-stressed parents. The compensatory photosynthetic responses of offspring ramets connected to stressed parents revealed an increasing trend as the experiment progressed. Morphological and physiological plasticity of F. tikoua in response to heterogeneous water resources allow them to adapt to karst habitats and be suitable candidates for vegetation restoration projects.